Willdenowia 40 – 2010 205

Siti Nurfazilah bt Abdul Rahman1, Ahmad Sofiman Othman1, Mohd Fahmi bin Abu
Bakar2 & Peter C. Boyce1*

Studies on Hanguana (Commelinales, Hanguanaceae) for Sunda II: Five new forest
species from Peninsular Malaysia and recircumscription of Hanguana malayana

Abstract
Siti Nurfazilah A. R., Ahmad Sofiman O., Mohd Fahmi A. B. & Boyce P. C.: Studies on Hanguana (Commelinales,
Hanguanaceae) for Sunda II: Five new forest species from Peninsular Malaysia and recircumscription of Hanguana
malayana. – Willdenowia 40: 205 –219. – Online ISSN 1868-6397; © 2010 BGBM Berlin-Dahlem.
doi:10.3372/wi.40.40206 (available via http://dx.doi.org/)
Field work in Peninsular Malaysia as part of the first author’s research into relationships and evolutionary polarity be-
tween Hanguana malayana, a widespread freshwater helophyte, and the numerous and mostly undescribed Sundaic
forest species has to date resulted in the collection of five distinctive novel forest-dwelling Hanguana species. These
are described here and figured in colour. Increased understanding of these and other forest species of Hanguana has
enabled a concise delimitation of the taxonomically long-obfuscated colonial helophytic H. malayana, and this is
here presented, and the species figured. The current taxonomy of Hanguana is summarised, and a key to the so-far
described Peninsular Malaysian species is provided.
Additional key words: monocots, taxonomy, mesophytes, helophytes

Introduction
Although the taxonomy of Hanguana (Hanguanaceae)
in Sunda remains beset with considerable confusion,
field work in Malaysia is beginning to permit better un-
derstanding of the many taxonomic novelties and through
this gradually establish a stable taxonomic platform from
which to begin phylogenetic investigations of the genus
Hanguana (Nurfazilah & al., in press).
Aside from making a start to bring taxonomic clar-
ity to forest species of Hanguana, ongoing field work in
Peninsular Malaysia has facilitated correction of several
morphological misconceptions, ecological inexactitudes,
and taxonomic and nomenclatural problems associated
with Hanguana malayana (Jack) Merr.
Results
Taxonomy of Hanguana
Hanguana Blume, Enum. Pl. Javae: 15. 1827.
Type: Hanguana kassintu Blume
=  Susum Blume ex Roem. & Schult., Syst. Veg. 7(2):
95. 1830. – Type: Susum anthelminthicum Blume ex
Roem. & Schult.
=  Veratronia Miq., Fl. Ned. Ind. 3: 553. 1859. – Type:
Veratronia malayana (Jack) Miq.
References. — Backer 1924: 3, 1951: 248 – 250; Airy Shaw
1965: 260 – 261; Rudall & al. 1999: 311 – 330; Tillich &
Sill 1999: 215 – 238; Dassanayake 2000: 214 – 215.
1. Hanguana bakoensis Siti Nurfazilah, Sofiman Othman
& P. C. Boyce, in press. – Holotype: Malaysia, Sarawak,
Bahagian Kuching, Bako N. P., Lintang Trail, 27.5.2007,
Nadiah I., Malcom D., Army K. & al. S.100599 (SAR!;
isotype: KEP, n.v.)

The changes and novel additions proposed in this paper 2. Hanguana bogneri H.-J. Tillich & E. Sill in Sendt-
enable the current taxonomy of Hanguana, comprising nera 6: 216. 1999. – Holotype: Malaysia, Sarawak, bei
10 species, to be summarised as follows: der Orang-Utan-Station Semenggoh, J. Bogner 94/2211;

1 Pusat Pengajian Sains Kajihayat [School of Biological Sciences], Universiti Sains Malaysia (USM), 11800 Pulau Pinang, Ma-
laysia; *e-mail: phymatarum@gmail.com; hp:/mobile/cell +6 017 329 2609 (author for correspondence).
2 Department of Biotechnology, Faculty of Agriculture and Biotechnology, Universiti Darul Iman Malaysia (UDM), Malaysia
Kampus Kota, Jalan Sultan Mahmud, 20400 Kuala Terengganu, Malaysia.
206 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia
kultiviert im Botanischen Garten München; 21.10.1996
(M!).
3. Hanguana exultans Siti Nurfazilah, Mohd Fahmi,
Sofiman Othman & P. C. Boyce, present paper.
4. Hanguana kassintu Blume, Enum. Pl. Javae: 15. 1827
≡ Susum kassintu (Blume) Kurz in Flora 56: 224. 1873 ≡
Susum malayanum f. sylvatica Backer, Handb. Fl. Java 3:
3. 1924 ≡ Hanguana malayana subsp. kassintu (Blume)
Backer, Bekn. Fl. Java 10(212): 2. 1949. – Holotype: In-
donesia, Java “crescit in sylvis montium Seribu Javae in-
sulae”, Blume s.n. (L!; isotype: BO!).
Notes. — Backer (1951) followed Merrill (1915) in
assigning Hanguana kassintu as a synonym of H. ma­
layana, but provided no detailed discussion. In fact, it
is obvious from the protologue discussion (Blume 1827)
and from examination of the type collection that H. kas­
sintu is a forest species fully distinct from H. malayana.
Blume, who knew both taxa, was perfectly aware of this.
That H. kassinutu is distinct is also implied by Airy Shaw
(1978), who, under the synonymy of H. malayana, cited:
“H. kassintu auctt., pro parte, vix Bl. Enum. Pl. Javae: 15
(1827)”. H. kassintu and the other Javan species will be
the subject of another paper in this series.
5. Hanguana major Airy Shaw in Kew Bull. 35: 819.
1981. – Holotype: Malaysia, Sabah, Kinabalu, Mesi-
lau River, 1500  m [converted from feet on the label],
5.2.1964, Chew & Corner RSNB 4233 (K!).
Notes. — There are problems with the circumscription of
Hanguana major. The plate accompanying the descrip-
tion does not feature any type elements and in fact rep-
resents two additional and furthermore novel taxa, with
one (Chai S 34089) not cited in the material seen. As yet
we have not examined all the cited material; a paper deal-
ing with the taxonomy of H. major and the other species
inadvertently included in the original publication awaits
completion of this examination.
6. Hanguana malayana (Jack) Merr. in Philipp. J. Sci.,
C, 10: 3. 1915 ≡ Veratrum malayanum Jack, Malayan
Misc. 1(1): 25. 1820
=  Hanguana anthelminthica (Blume ex Roem. &
Schult.) Masam., Enum. Phan. Born.: 81. 1942 ≡ Su­
sum anthelminthicum Blume ex Roem. & Schult.,
Syst. Veg. 7(2): 1493. 1830
=  Hanguana aquatica Kaneh. in Trans. Nat. Hist. Soc.
Formosa 25: 8. 1935.
Note. — A full synonymy and revised circumscription is
provided in the present paper below.
7. Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofi-
man Othman & P. C. Boyce, present paper.
8. Hanguana pantiensis Siti Nurfazilah, Mohd Fahmi,
Sofiman Othman & P. C. Boyce, present paper.
9. Hanguana podzolicola Siti Nurfazilah, Mohd Fahmi,
Sofiman Othman & P. C. Boyce, present paper.
10. Hanguana stenopoda Siti Nurfazilah, Mohd Fahmi,
Sofiman Othman & P. C. Boyce, present paper.
Insertae sedis
Susum minus Miq., Fl. Ned. Ind., Eerste Bijv.: 598.
1861. – Holotype: Indonesia, Sumatera Tengah, Paria-
man (‘prov. Priaman’) (BO?, L?, not yet traced).
Notes. — Miquel’s protologue describes this plant in
some detail, comparing it against Blume’s Susum anthel­
minthicum (= Hanguana malayana). In particular he not-
ed the marked disparity in size between the inner larger
and outer smaller sepals, the sepals to be “laciniae” and
also the much smaller fruit (“baccis multo minoribus”).
The plant is also noted as being glabrous. In the contin-
ued absence of an authentic specimen it is impossible to
ascribe Miquel’s epithet. Although assuming Miquel’s
observation of laciniate tepals is correct, it is a feature
otherwise not recorded in any Hanguana with which we
are familiar. We are therefore in no doubt that it repre-
sents a distinct, almost certainly forest-dwelling taxon.
Key to peninsular Malaysian Hanguana species
1. Stoloniferous colonial helophytes  . . . . . . . . . . . . .   2
– Clumping mesophytes lacking stolons  . . . . . . . . .   3
2. Leaves stiffly erect, acute; fruits ellipsoid, ripening
glossy purple-red; stigma lobes flat, connate at base,
the whole 3 – 4  mm diam. and almost obscuring the
end of the fruit. Plants of open situations along muddy
banks of large rivers, margins of freshwater bodies,
and of freshwater swamp forest  . . . . .   H. malayana
– Leaves arching, long-attenuate; fruits globose, ripen-
ing semiglossy black; stigma lobes small, separate,
erect, pointed. Plants of shaded peatswamp mires . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .   H. nitens
3. Stigma inserted obliquely  . . . . . . . . . . . . . . . . . . .   4
– Stigma terminal  . . . . . . . . . . . . . . . . . . . . . . . . . . .   5
4. Fertile portion of infructescence not exceeding leaves,
panicle dense, branches of the partial inflorescences
ascending in fruit; plants sessile even in old age  . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .   H. pantiensis
– Fertile portion of infructescence far exceeding leaves,
panicle very open, branches of the partial inflores-
cences spreading and forming regular tiers; plants de-
veloping an erect, leafless stem up to 1.5 m tall  . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .   H. podzolicola
5. Infructescence with caducous foliaceous bracts; par-
tial inflorescences each with 2 or rarely 3 branches,
spreading in fruit; fruits globose with a briefly stipi-
Willdenowia 40 – 2010
tate stigma, lobes connate basally, deep chocolate
brown; fruit ripening white with conspicuous black
speckles; old plants developing a short (to c. 25 cm)
slender, leafless stem. Plants of well-drained slopes
and low ridges in lowland humid, moist mixed dipte-
rocarp forest on yellow clay soils  . . .   H. stenopoda
– Infructescence with persistent foliaceous bracts;
partial inflorescences each with 4 – 5 branches, these
rather sharply ascending in fruit; fruits ventrally gib-
bose-ellipsoid, stigma sessile, comprising 3 free or-
ange brown lobes; fruit ripening pale yellow without
conspicuous black speckles. Plants of low-lying wet
podzols in peatforest  . . . . . . . . . . . . . .   H. exultans
New forest species from Peninsular Malaysia
Hanguana exultans Siti Nurfazilah, Mohd Fahmi, Sofi-
man Othman & P. C. Boyce, sp. nov.
Holotypus: Malaysia, Johor Bahru, Kota Tinggi, Hutan Li-
pur Panti, 1°48'07.7''N, 103°57'20.2''E, 40 m, 20.4.2010,
Siti Nurfazilah bt Abdul Rahman, P. C. Boyce & Ooi Im
Hin HA-55 (KEP!).
Hanguanae stenopodae maxime similis, facile inflores-
centiis partialibus omnibus ramis 4 vel 5, his fructu satis
valde ascendentibus, tepalis interioribus fere diaphanis,
infrutescentiis bracteis persistentibus foliaceis, fructibus
ventraliter gibboso-ellipsoideis, maturatione dilute luteis
sine punctis conspicuis nigris, stigmate sessili, lobis 3 sep-
aratis (non connatis) distinguenda.
Solitary medium-sized, herbaceous, dioecious mesophyte
to c. 1 m tall. Leaves up to 1.8 m long; up to 15 togeth-
er, semi-erect, tips arching, somewhat flocculose abaxi-
ally, soon glabrescent, bases imbricate; leaf blade up to
90 × 15  cm, elongate-elliptic, base long-decurrent, tip
rather attenuate; glossy deep green when fresh, drying
dull olive-green abaxially, silver-green adaxially; pseu­
dopetiole up to 80 cm long, 1 cm wide, accounting for
almost 1/2 the entire leaf length, shallowly channelled,
the margins somewhat sharp, longitudinally folded in-
wards, petiole very pronounced, c. 24 cm long, petiolar
sheath with margins wide, glossy erose-marcescent, dry-
ing deep brown; midrib somewhat prominently round-
raised abaxially, drying shallowly channelled for c. 2/3 of
the blade, flush distally adaxially and all other venation
prominent minutely tessellate-striate in most specimens.
Female and male inflorescences not observed, although,
based on fruiting material and observations of inflores-
cence architecture, almost certainly erect at anthesis. In­
fructescence erect, thyrsoid-paniculate, not exceeding the
leaves, comprising up to 6 partial alternate-secund, thyr-
soid infructescences plus a terminal spike; peduncle and
scape together up to 60 cm tall, conspicuously very pale
brown floccose, especially the scape, dark green when
fresh, drying dark brown, visible portion of peduncle up
to 9  cm long; bract marking start of scape foliaceous,
207
sterile, persistent, broadly lanceolate, 52 × 10.5 cm, base
clawed, apex long-attenuate; subtending bract of partial
infructescences similar to that marking the start of the
scape, diminishing in size distally along the infructes-
cence, the largest c. 19.5 × 2 cm, the smallest 15 × 3 mm,
all infructescence bracts drying pale straw-coloured; par­
tial infructescences each comprising up to 7 branches,
often fewer, arising simultaneously from the axil of the
subtending bract, median branches c. 9 cm long, lateral
branches up to 5 cm long, both 1 – 1.5 mm wide, somewhat
erect. Female flowers scattered, mainly solitary, sessile, all
with an associated minute bracteole; perianth of 6 trans-
parent, very pale green tepals, outermost c. 1.9 × 1.1 mm,
ovate, inner tepals c. 2.1 × 1.9 mm, ovate, all tepals clasp-
ing the base of the fruit. Ripe fruits ventrally gibbose-
ellipsoid, pale yellow; stigma sessile when fresh, drying
conspicuously raised, c. 1.2 mm diam., comprising 3 free
orange-brown lobes. Seeds not observed. – Fig. 1.
Ecology. — Low-lying wet (but not swampy) podzols
under closed-canopy lowland humid moist peatforest.
Notes. — Hanguana exultans is most similar to H. ste­no­
poda although readily distinguished by the nearly trans-
parent inner tepals, infructescence with persistent folia-
ceous bracts, partial inflorescences each with 4 – 5 branches
rather sharply ascending in fruit, ventrally gibbose-ellip-
soid fruits with a sessile stigma comprising 3 separate (not
connate) orange brown lobes and the fruit ripening pale
yellow without conspicuous black speckles.
Older plants of Hanguana exultans do not develop
the slender, short, naked stem typical for H. stenopoda.
There are also ecological differences: H. exultans is a
plant of low-lying wet podzols in peatforest, whereas H.
steno­poda is restricted to well-drained slopes and low
ridges in lowland humid, moist mixed dipterocarp forest
on yellow clay soils.
Etymology. — Latin “exultans” = boastful, vainglorious,
in rather fanciful allusion to this species remaining aloof
from the more abundant and parapatric Hanguana pan­
tiensis.
Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman
Othman & P. C. Boyce, sp. nov.
Holotypus: Malaysia, Johor Bahru, Mersing, Hutan
Simpanan Lenggor, 2°15'72.7''N, 103°43'76.7''E, 55  m,
18.4.2010, Siti Nurfazilah bt Abdul Rahman, P. C. Boyce
& Ooi Im Hin HA-48 (KEP!).
Hanguanae malayanae superficialiter habitu stolonifero
coloniali similis, sed facile foliis arcuatis (non rigide
erectis) graviter nitide viridibus, lamina conspicue longa
(ad 2/3 – 3/4 folii longitudine) pseudopetiolata leniter plica-
ta, apice longe attenuato, fructibus globosis maturatione
seminitide nigris, stigmatis lobis separatis erectis acutis
distinguenda.
208 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia
Medium-sized, glabrous, dioecious stoloniferous helo-
phyte to c.  1  m tall; stem terete, rhizomatous with the
terminal portion ascending, up c.  1.5  cm diam., sub-
woody, the older portions clothed in dense fibrous de-
graded leaf bases; stolons up to 45  cm long (although
usually less), c. 1.5 cm diam., semi-erect or creeping, or
burrowing through liquid peat, enveloped by appressed
petiolar sheaths, these deep green and foliaceous towards
the stolon active tip, older leaf portions turning glossy
chestnut brown and eventually partially or completely
decaying into fibres. Leaves up to 1.35  m long, up to
16 together, long-pseudopetiolate, ± erect, innovations
sometimes very sparsely flocculose abaxially along the
midrib, soon glabrescent, with age entirely glabrous; leaf
blade 45 – 100 × 5 – 8  cm, narrowly lanceolate, leathery,
dark shining green when fresh, drying dull olive-green,
the base decurrent on the pseudopetiole, tip long-atten-
uate; pseudopetiole accounting for c. 2/3 – 3/4 of the leaf
length, ± V-shaped in cross-section, margins sharp, peti-
olar sheath margins drying papery; midrib acutely raised
abaxially, flush to slightly impressed adaxially, blade
irregularly but prominently plicate when fresh, densely
longitudinally veined, with numerous close-set very
fine cross-veinlets. Female and male inflorescences a
moderately stout pedunculate panicle, erect at anthesis,
subtended by a fully developed foliage leaf, with up to
15 pa­tently branched thyrsoid or spicate partial inflores-
cences plus a terminal spike; bract subtending proximal
partial inflorescences very narrowly triangular, folia-
ceous, 5 – 24 cm long, 0.5 – 2 cm wide, bract subtending
distal-most partial inflorescences small to minute, ovate,
apiculate; median branches longer than lateral branches,
up to 15 cm long and c. 3 mm wide, rectangular in cross
section at the base, angle further up, lateral branches ap-
proximately 3.5 cm long; terminal spike c. 9 cm long, all
branches weakly ascending; peduncle and scape together
up to 80  cm tall, lower part of peduncle up to 1.5  cm
diam., the whole inflorescence weakly flocculose, gla-
brescent or nearly so in part, pale green with areas of
purple-brown speckles and spots when fresh, drying deep
brown with remaining hairs pale brown, visible portion
of peduncle up to 30  cm long; bract marking onset of
scape large, foliaceous, sterile, c. 60 cm long, 9 cm wide,
narrowly triangular, base clasping, tip long-attenuate.
Male flowers not observed. Female flowers ± distant,
mostly solitary, occasionally in small clusters, especially
towards the tips of branches, sessile with a broad base
in the axil of a short, broad bract; tepals shortly con-
nate at the base, green, outer 3 tepals c. 1 × 2.5 mm, in-
ner tepals c. 2 × 2.5 mm; staminodes not observed; ovary
broadly ovoid-globose, pale green, ripening through
yellow-green to black; stigma sessile, 3-lobed, c. 2 mm
diam., lobes separate, erect, acute-pointed, matte black.
Infructescence erect. Ripe fruit globose, c. 4 mm diam.,
semiglossy black, stigma remains matte black. Seeds
hemispherical, deeply excavated, with the pit margins in-
curved, c. 3 mm diam. – Fig. 2.
Distribution. — Malaysia, Johor Bharu.
Ecology. — Blackwater mires over saturated deep peat
along margins of closed-canopy lowland humid moist
peatswamp forest at altitudes of 40 – 50 m.
Notes. — Hanguana nitens by its stoloniferous, colonial
habit is superficially similar to H. malayana, but may be
readily distinguished by arching (not stiffly erect), deep
glossy green leaves, with the blade conspicuously long-
pseudopetiolate (up to 2/3 – 3/4 of the leaf length) and
weakly plicate with a long-attenuate tip, and globose fruit
ripening semiglossy black with separate, erect, pointed
stigma lobes.
So far, Hanguana nitens is known from three sites.
At the type locality it is abundant, occurring as several
moderately extensive female stands, and a single isolated
male population. The second known locality, Hutan Sim-
pan Panti, consists of a much smaller single population
of female plants. The third site, at Hutan Lipur Panti, is a
relict, very much depleted unsexed population critically
threatened by an adjacent building development.
Etymology. — Latin “nitens” = shining, polished, in allu-
sion to the remarkably lustrous leaf blade.
Other material seen. — Malaysia: Johor Bahru, Kota
Tinggi, Hutan Simpan Panti, 1°52'22.6''N, 103°54'75.5''E,
19.4.2010, Siti Nurfazilah bt Abdul Rahman, P. C. Boyce
& Ooi Im Hin HA-51 (KEP); Johor Bahru, Kota Tinggi,
Hutan Lipur Panti, 1°48'07.7''N, 103°57'20.2''E, 20.4.
2010, Siti Nurfazilah bt Abdul Rahman, P. C. Boyce &
Ooi Im Hin HA-57 (KEP).
Hanguana pantiensis Siti Nurfazilah, Mohd Fahmi, So­
fiman Othman & P. C. Boyce, sp. nov.
Holotypus: Malaysia, Johor Bahru, Kota Tinggi, Hutan
Lipur Panti, 1°48'07.7'', 103°57'20.2'', 40 m, 19.4.2010,
Siti Nurfazilah bte Abdul Rahman, P. C. Boyce & Ooi Im
Hin HA-56 (KEP!).
Hanguanae podzolicola maxime similis, aetate sine cul-
mo longo efoliato, pedunculo breviore, inflorescentiae
parte fertili foliis non excedenti, paniculae structura con-
fertiore, inflorescentiae partialis ramis fructu ascendenti-
bus (non effusis) differt.
Solitary robust, herbaceous, dioecious mesophyte to
c. 2 m tall. Leaves up to 1.6 m long, up to 16 together,
spreading, flocculose abaxially, the outermost leaves
arching with the tips touching the ground, bases imbri-
cate; leaf blade up to 84 × 13.5 cm, narrowly elliptic, base
decurrent, tip long-attenuate with a conspicuous apicule
to 5 mm, deep green semiglossy when fresh, drying dull
olive-green abaxially, silver-green adaxially; pseudopeti­
ole up to 15 cm long, 1 cm wide, accounting for c. 1/5 of
the entire leaf length, shallowly channelled, the margins
Willdenowia 40 – 2010
somewhat sharp, longitudinally folded inwards, lower-
most part of petiolar sheath with margins erose-marces-
cent; midrib prominently round-raised abaxially, espe-
cially in the lower middle part of the leaf blade, impressed
adaxially, drying flush adaxially and all other venation
prominent minutely tessellate-striate in most specimens.
Female and male inflorescences not observed, although,
based on fruiting material and observations of inflores-
cence architecture, almost certainly erect at anthesis. In­
fructescence erect, dense-paniculate, not exceeding the
leaves, comprising up to 13 partial, whorled, thyrsoid or
more rarely spicate infructescences plus a terminal spike;
peduncle and scape together up to 70 cm tall, conspicu-
ously very pale brown flocculose, especially the scape,
dark green when fresh, drying dark brown, visible por-
tion of peduncle up to 20 cm long; bract seemingly mark-
ing start of scape foliaceous, sterile, lanceolate-elliptic,
soon falling (but more investigation needed); subtending
bract of partial infructescences similar to that marking
the start of the scape, diminishing in size distally along
the infructescence, the largest c.  48 × 10  cm, the small-
est 18 × 3  mm; partial infructescences each comprising
up to 12 branches arising simultaneously from the axil of
the subtending bract, median and lateral branches almost
equal in length, up to 14 cm long, 1 – 2 mm wide, mark-
edly erect. Female flowers scattered, mainly solitary, ses-
sile, all with an associated minute bracteole; perianth of
6 opaque, green tepals, outermost c.  1.5 × 2 mm, ovate,
weakly concave, inner tepals c. 2 × 2.5 mm, ovate, all te-
pals clasping the base of the fruit. Ripe fruit pink, with
minute raised yellow speckles, oblique-globose, c. 5 mm
diam., ripening glossy pink; stigma 3-lobed, lobes not
connate, c.  1  mm, raised, dark brown. Seeds not ob-
served. – Fig. 3.
Distribution. — Only known from the type locality,
where it is abundant.
Ecology. — Well-drained flat and slightly undulating
areas of lowland closed canopy humid moist peatforest.
40 – 60 m.
Notes. — Hanguana pantiensis is one of a group of spe-
cies, all novel, in which early in development the ovary
bends longitudinally to produce a fruit with the stigma
obliquely to sublaterally inserted. H. pantiensis is most
similar to H. podzolicola, differing by the much shorter
peduncle, with the fertile portion of the inflorescence not
exceeding the leaves, the denser panicle structure, with
the branches of the partial inflorescence ascending (not
spreading) in fruit and by not developing a tall, leafless
stem with age.
Hanguana pantiensis is abundant at the type local-
ity, forming dense pure stands. At the time of collection
several female plants were fruiting, and numerous plants
with spent inflorescences suggested that male plants were
common.
209
Etymology. — Type locality “Panti” + Latin “ensis” = ori­
ginating from, referring to the type and only known lo-
cality.
Hanguana podzolicola Siti Nurfazilah, Mohd Fahmi,
Sofiman Othman & P. C. Boyce, sp. nov.
Holotypus: Malaysia, Johor Bahru, Mersing, Hutan
Simpanan Lenggor, 2°15'72.7''N, 103°43'76.7''E, 55 m,
18.4.2010, Siti Nurfazilah bt Abdul Rahman, P. C. Boyce
& Ooi Im Hin HA-50 (KEP!).
Hanguanae pantiensi maxime similis in stigmatis inser-
tione obliqua, sed paniculae structura plurime aperta,
inflorescentiae partialis fructu ramis effusis (non valde
ascendentibus), praeterea individuis vetioribus caule
erecto efoliato evoluerentibus distinguenda.
Solitary robust, herbaceous, dioecious mesophyte to
c. 2 m tall, stem erect, with age becoming leafless to 1.5 m
with a terminal crown of leaves. Leaves up to 1.5 m long,
up to 15 together, spreading, bases imbricate; leaf blade
up to 80 × 13 cm, narrowly elliptic, base decurrent, tip
long-attenuate with a conspicuous apicule to 7 mm, stiffly
chartaceous and deep green when fresh, drying somewhat
softer and dull olive-green abaxially, silver-green adaxi-
ally; pseudopetiole c. 45 cm long, 2 cm wide, account-
ing for up to 1/3 of the entire leaf length, shallowly chan-
nelled, the margins very sharp, especially about half way
along the length; petiolar sheath margins hyaline when
fresh, drying medium-brown and ultimately marcescent;
midrib pronouncedly round-raised abaxially, channelled
adaxially; lesser venation very obscure when fresh, dry-
ing closely and conspicuously longitudinally-raised stri-
ate with pronouncedly reticulate lesser venation, with the
marginal veins drying darker adaxially and overtopping
the leaf blade to form the apicule. Female and male in­
florescences not observed, although, based on fruiting
material and observations of inflorescence architecture,
almost certainly erect at anthesis. Infructescence soli-
tary, erect, comprising up to 11 partial, whorled, thyr­
soid or more rarely spicate infructescences plus a termi-
nal spike; peduncle and scape together up to 80 cm tall,
conspicuously very pale brown flocculose, green when
fresh, drying dark brown, visible portion of peduncle up
to 30  cm long; bract seemingly marking start of scape
foliaceous, sterile, lanceolate-elliptic, soon falling (but
more investigation needed); bract subtending partial
infructescences similar to that marking the start of the
scape, diminishing in size distally along the infructes-
cence, the largest c. 41 × 5.5 mm, the smallest 25 × 5 mm;
partial infructescences each comprising of 9 branches,
branches arising simultaneously from the axil of the sub-
tending bract, median and lateral branches almost equal
in length, 10 – 18  cm long, 1 – 2  mm wide, somewhat
flattened. Female flowers scattered, mainly solitary, oc-
casionally in groups of 2, sessile, all with an associated
minute bracteole; perianth of 6 opaque, green tepals, out-
210 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia
ermost c. 1 × 1.8 mm, ovate, weakly concave, inner tepals
c. 2 × 2 mm, ovate, all clasping the base of the fruit. Ripe
fruit pale yellow with minute pink speckles, oblique-glo-
bose, c. 5 mm diam., ripening glossy medium pink to ma-
genta; stigma 3-lobed, lobes connate basally, c. 1.3 mm,
raised, matte deep brown. Seeds not observed. – Fig. 4.
Distribution. — Malaysia, Johor Bahru.
Ecology. — Raised podzols in closed canopy lowland
humid moist peatforest, occasionally on raised podzols
in seasonally swampy peatforest. 40 – 60 m.
Notes. — Hanguana podzolicola closely resembles H.
pantiensis by the oblique insertion of stigma, but may be
distinguished by the much more open panicle structure,
partial inflorescence branches spreading (not sharply as-
cending) in fruit, and by older individuals developing an
erect, leafless stem up to 1.5 m tall.
Etymology. — Latin “cola” (derived from “incola”, the
inhabitant, dweller) + “podzol”, a raised peat deposit. Re-
ferring to the restricted ecology of this species.
Other material seen. — Malaysia, Johor Bahru, Kota
Tinggi, Hutan Lipur Panti, 1°48'07.7''N, 103°57'20.2''E,
19.4.2010, Siti Nurfazilah bt Abdul Rahman, P. C. Boyce
& Ooi Im Hin HA-53 (KEP).
Hanguana stenopoda Siti Nurfazilah, Mohd Fahmi,
Sofi­man Othman & P. C. Boyce, sp. nov.
Holotypus: Malaysia, Pahang, Rompin, Taman Negeri
Endau Rompin, 2°37'37.2''N, 103°19'83.3''E, 12.5.2010,
Siti Nurfazilah bt Abdul Rahman, Mohd Fahmi Bin Abu
Bakar & P. C. Boyce HA-60 (KEP!).
Hanguanae exul tanti similis, inflorescentiis partialibus
omnibus ramis 2 raro 3, his fructu effusis, tepalis interiori-
bus opace viridibus, infructescentiis bracteis caducis fo-
liaceis, fructibus maturitate albis punctis conspicue nigris,
stigmate breviter stipitato lobis basaliter connatis graviter
badiis differt. Maturitate caule brevi gracili efoliato.
Solitary, herbaceous, dioecious mesophyte to c. 1 m tall,
developing a short (up to 25  cm), slender leafless stem
with age. Leaves up to 1 m long, up to 15 together, ini-
tially semi-erect, older (outermost) leaves spreading, the
longest leaves arching with the tips touching the ground,
bases imbricate; leaf blade up to 51.5 × 9  cm, elliptic,
thinly leathery, glossy, bright green when fresh, drying
thinly chartaceous and pale straw-coloured; pseudopet­
iole 35–44  cm long, c.  1  cm wide, accounting for just
under 1/2 of the entire leaf length, when fresh somewhat
channelled, with the margins very sharp, drying strongly
longitudinally folded; lowermost part of petiolar sheath
c.  16  cm long, margins rather wide, erose-marcescent
leaf tip long-attenuate, tipped with a 1 cm tubule; mid­
rib prominent proximally, somewhat semiterete, raised
abaxially, especially in the lower part of the leaf blade,
distally flush, somewhat impressed adaxially until the
mid-point, thereafter flush, other venation striate, ob-
scure when fresh, drying somewhat prominent, with all
minor venation minutely tessellate. Female and male
inflorescences not observed, although, based on fruit-
ing material, almost certainly erect at anthesis. Infruct­
escence erect, comprising 4 partial, thyr­soid partial in-
fructescences plus a terminal spike, subtended by a fully
developed foliage leaf; peduncle and scape together up
to 62 cm tall, greyish flocculose, dark brown-red, visible
portion of peduncle up to 22.5 cm long, 5 mm wide; bract
marking start of scape foliaceous, sterile, narrowly ellip-
tic, up to 25 × 6.5  cm, base clawed, tip attenuate; bract
subtending partial infructescences linear-triangular, up to
65 × 6 mm, diminishing in size distally along the infruct-
escence, the smallest c. 7 × 2 mm; partial infructescences
each comprising of 3 branches, the terminal one a single
spike, branches arising simultaneously from the axil of
the subtending bract, median branch usually longer than
lateral branches, 5 – 7 cm long, c. 1.5 mm wide, weakly
angled, lateral branches approximately 2/3 of the length
of median branch, although branches subequal in distal-
most partial infructescence. Female flowers mainly in
scattered groups of 2 to 3, lowermost flowers of each
branch occasionally solitary, all mainly sessile, very oc-
casionally pedicellate to c.  0.5  mm, all with an associ-
ated minute bracteole; perianth of 6 tepals, outer tepals
1.2 × 1.9 mm, ovate, inner tepals c. 2.5 × 2.5 mm, ovate,
all clasping fruits in fresh material and medium green,
outer with a narrow dark red margin, inner only red at the
tip. Ripe fruit compressed-globose, c. 5 × 4.2 mm, glossy
white with conspicuous black speckles; stigma 3-lobed,
lobes connate to form a clover-leaf, c. 1.9 mm diam., very
shortly stipitate, matte black. Seeds not observed. – Fig. 5.
Distribution. — Malaysia, Pahang, known only from the
type locality.
Ecology. — Lowland humid, moist mixed dipterocarp
forest on yellow clay soils with a moderate leaf litter layer
at altitudes of 60 – 125 m.
Notes. — Hanguana stenopoda resembles H. exultans,
differing by the opaque green inner tepals, infructes-
cence with caducous foliaceous bracts, partial inflores-
cences each with 2 or rarely 3 branches, these spreading
in fruit, globose fruits with a briefly stipitate stigma, in
which the lobes are connate basally and deep chocolate
brown, and the fruit ripening white with conspicuous
black speckles. In age, H. stenopoda develops a short,
slender, leafless stem.
Hanguana stenopoda is restricted to well-drained
slopes and low ridges in lowland humid, moist mixed dip­
terocarp forest on yellow clay soils with a moderate leaf
litter layer.
Willdenowia 40 – 2010
Etymology. — Greek FJg<@H (stenos) = slender, and Lat-
in podos = a foot, referring to the short, slender stem that
develops in older individuals.
Other material seen. — Malaysia, Pahang, Temerloh,
Gunung Benom, Krau Wildlife Sanctuary 3°49'29.8''N,
102°12'84.6''E, 125, 13.5.2010, Siti Nurfazilah bt Abdul
Rahman, Mohd Fahmi Bin Abu Bakar & P. C. Boyce HA-
61 (KEP).
Hanguana malayana recircumscribed
Hanguana malayana (Jack) Merr. in Philipp. J. Sci.,
C, 10: 3. 1915 ≡ Veratrum malayanum Jack, Malayan
Misc. 1(1): 25. 1820 ≡ Veratronia malayana (Jack) Miq.,
Fl. Ned. Ind. 3: 553. 1859 ≡ Susum malayanum (Jack)
Planch. ex Hook. f., Fl. Brit. India 6: 391 1892. – Holo-
type: Malaysia, Pulau Pinang [“Poeloe Pinang”], Jack s.n.
(not traced). – Epitype (designated here): Wallich EIC
3717 [K-WAL!].
=  Hanguana anthelminthica (Blume ex Roem. &
Schult.) Masam., Enum. Phan. Born.: 81. 1942 ≡ Su­
sum anthelminthicum Blume ex Roem. & Schult., Syst.
Veg. 7(2): 1493. 1830 ≡ Susum malayanum f. aquati­
ca Backer, Handb. Fl. Java 3: 3 (1924) ≡ Hanguana
malayana subsp. anthelminthica (Blume ex Roem.
& Schult.) Backer, Bekn. Fl. Java 10(212): 2. 1949
≡ Hanguana malayana var. anthelminthica (Blume
ex Roem. & Schult.) Bakh. in Blumea 6: 399 1950. –
Holo­type: Indonesia, Java “in paludibus circa Buiten-
zorg”, Blume s.n. (L!; isotype: BO!).
=  Hanguana aquatica Kaneh. in Trans. Nat. Hist. Soc.
Formosa 25: 8. 1935. – Holotype: Caroline Islands,
Palau, Almonogni, Babeldaob, 13.4.1938, S. Hatusima
4866 (FU!).
References. — Ridley 1907: 131 – 133, 1924: 369; Backer
1951: 248 – 250; Airy Shaw 1975: 1 – 50.
Large to massive colonial stoloniferous dioecious helo-
phytes to c.  3  m tall, exceptionally reaching 4  m; stem
terete, robustly rhizomatous with the terminal portion as-
cending, up c. 15 cm diam., although usually somewhat
less, spongy, the older portions clothed in dense fibrous
degraded leaf bases; stolons up to 2.5  m long (usually
less), c. 2 cm diam., creeping or burrowing through li­quid
mud, or floating on the surface of water, enveloped by
appressed sheath, these foliaceous towards the stolon ac-
tive tip, which finally dissolve into fibres. Leaves excep-
tionally up to 3.5 m long, more usually about 1.5 – 2 m,
up to 20 together, stiffly erect, occasionally fleetingly
flocculose abaxially when young, otherwise glabrous;
lower leaves briefly pseudopetiolate, higher leaves (of
flowering plants) remote, smaller; on shorter petioles,
pedunculate and scapose bracts sessile or subsessile with
a broad base, small, passing into fertile bracts; lower
leaf sheaths long and broad, stem clasping, deeply split
211
on the anterior side, gradually narrowing into the peti-
ole; leaf blade ­(20 –)45 – 200(– 300) × 4 – 15  cm, lanceo-
late, somewhat spongy-leathery, medium to dark green,
sublustrous when fresh, drying spongy-chartaceous and
medium straw-coloured, base acute to decurrent on the
pseudopetiole, tip acute; pseudopetiole accounting for
c. 1/5 of the leaf length, ± flat to shallowly V-shaped, mar-
gins sharp, lowermost part of petiolar sheath margins
later marcescent; midrib rather thick, blade densely lon-
gitudinally veined, with numerous close-set thin cross-
veinlets, between the longitudinal veins very densely and
finely longitudinally striate, not or only very obscurely
plicate. Female and male inflorescences a stout pedun-
culate panicle erect at anthesis and subtended by a fully
developed foliage leaf, with up to 10 whorls of 1 – 10 ×
patently branched thyrsoid or spicate partial inflores-
cences plus a terminal spike; branches arising simulta-
neously from the axil of the subtending bract, median
branches usually longer than lateral branches, although
branches subequal in distal-most units; partial male in­
florescences 30.5 cm long, c. 4 mm wide, weakly angled,
lateral branches approximately 29  cm long, c.  2.1  mm
wide, thinner and longer than those of female inflores-
cences, rather distinctly yellowish, with a greater number
of flowers, branches initially ascending, later drooping
at post-anthesis prior to withering; partial female inflo­
rescences stout and short, 18  cm long, c.  4  mm wide,
weakly angled, lateral branches approximately 11  cm
long and c. 3 mm wide, green, ascending; peduncle and
scape together often more than 2  m tall, lower part of
peduncle up to 3 cm diam., the whole weakly to rather
densely flocculose, soon glabrescent or nearly so, medi-
um green, visible portion of peduncle up to 50 cm long;
bract marking onset of scape large, foliaceous, fertile, or
sterile, broadly lanceolate-elliptic, up to 20 cm long, base
clasping, tip long-acute. Flowers of both sexes ± distant,
either solitary or in small clusters, sessile with a broad
base in the axil of a short, broad bract and tepals shortly
connate at the base, green or yellowish or the inner tepal
dotted red. Male flowers with 3 outer tepals c. 1 mm long
and 3 inner tepals c. 1.5 cm long, fornicate; stamens 6, on
the base of the perianth, about as long as the inner tepals;
filaments filiform from a broader base, c.  2  mm long;
anthers small, c. 1 mm, with longitudinal slits, inserted
in a basal cleft; pistillode small, stigmas 3, erect, shortly
clavate. Female flowers with outer 3 tepals c. 2 mm long
and inner tepals c. 3 mm long; staminodes 6, c. 0.5 mm
long, inserted on the base of the perianth, those oppo-
site the outer sepals minute, narrowly triangular, 3 others
much longer and broader, rounded, dorsally compressed;
ovary broadly ovoid-globose, stigma sessile, deeply di-
vided into 3 spreading broadish short arms. Infructes­
cence erect, comprising up to 10 partial infructescences
each subtended by a semipersistent bract up to 20  cm
long, although these diminishing in size distally along
the infructescence, the smallest c.  10 × 2  cm. Ripe fruit
dark shining purple-red, ellipsoid, c.  1 × 5  mm, stigma
212 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia
3-lobed, lobes connate to form a clover-leaf almost ob-
scuring the end of the ovary, c. 3 mm diam., matte black.
Seeds not observed. – Fig. 6 – 7.
Distribution. — Equatorial tropics from Sri Lanka to
western Micronesia (Palau), south as far as northern Aus-
tralia and north to the Philippines (Luzon). Absent from
the area between the Wallace and Webber lines.
Ecology. — Plants of open lowland situations along mud-
dy banks of large rivers, margins of freshwater bodies,
and of freshwater swamp forest.
Notes. — Nurfazilah & al. (in press) have highlighted
that the Flora Malesiana account for Hanguana (Backer
1951) contains numerous misconceptions with the result
that the circumscription of H. malayana has been subject
to considerable and quite unnecessary confusion. Field
observations in Malaysia leave us in no doubt that H. ma­
layana is both morphologically and ecologically clearly
circumscribed, as defined in the above key.
It is interesting to note that Ridley (1907, 1924) was
clearly aware that the circumscription of Hanguana in
Peninsular Malaysia was in need of critical study. Unfor-
tunately, Ridley (1907) unaccountably applied the name
Susum malaynanum to the forest species (in the broad
sense), while for the open habitat helophyte he used the
(now synonymous) S. anthelminthica.
Backer (1951) stated the fruit to be shining red. This
is incorrect; the ripe fruit is dark shining purple-red, as
stated by Dassanayake & al. (1999).
Diligent searches of relevant herbaria have failed
to turn up Jack’s collection of Veratrum malayanum [≡
Hanguana malayana]. As alluded by Steenis-Kruseman
(2006) there are difficulties and obscurities associated
with Jack’s herbarium, not least because “an important
part of his botanical notes and collections was lost by the
fire of the ship ‘Fame’ on which Raffles embarked for
Europe in 1824”. It might be argued that the type is lost
and thus neotypification is justifiable. However, given that
Jack material periodically turns up in ‘expected’ places
(e.g. Cowan (1954) “methodically searched the Herb. Ed-
inburgh [E] for Jack specimens and found 64 in all (incl.
1 dupl.). No record of how and when they were acquired
was found …”), we are reluctant to take this step. As
a compromise, we have opted to epitypify Wallich EIC
3717, a Pinang collection as was Jack’s, and beyond any
doubt well-representative of the species.
Acknowledgements
Fieldwork in Malaysia was supported by a Research Uni-
versity Grant from the Universiti Sains Malaysia, Pen-
ang. Thanks go to J. F. Veldkamp (Leiden) for translation
of the Latin diagnoses.
References
Airy Shaw H. K. 1965: Diagnoses of new families, new
names, etc., for the seventh edition of Willis’s ‘Diction-
ary’. – Kew Bull. 18: 249 – 273. [CrossRef]
Airy Shaw H. K. 1978: Three interesting plants from the
Northern Territory of Australia (Thymelaeaceae, Fla­
courtiaceae & Hanguanaceae). – Kew Bull. 33: 1 – 5.
Airy Shaw H. K. 1981: A new species of Hanguana from
Borneo. – Kew Bull. 35: 819 – 821. [CrossRef]
Backer C. A. 1924: Handboek voor de flora van Java 3. –
Batavia: Ruygrock & Co.
Backer C. A. 1951: Hanguana. – Pp. 248 – 250 in: Steenis
C. G. G. J. van (ed.), Flora malesiana, ser. 1, 4(3). –
Djakarta: Noordhoff-Kolff.
Blume C. L. 1827: Enumeratio plantarum Javae et insu-
larum 1. – Leiden: Van Leeuwen.
Cowan J. M. 1954: Some information on the Menzies
and Jack collections in the herbarium, Royal Botanic
Garden, Edinburgh. – Notes Roy. Bot. Gard. Edin-
burgh 21: 219 – 227.
Dassanayake M. D. 2000: Hanguanaceae. – Pp. 214 – 215
in: Dassanayake M. D., Fosberg F. R. & Clayton W.
D. (ed.), A revised handbook to the flora of Ceylon
14. – Rotterdam: A. A. Balkema.
Merrill E. D. 1915: New or noteworthy Philippine plants
XI. – Philippine J. Sci., ser. C, 10: 1 – 84.
Ridley H. N. 1907: Materials for a Flora of the Malayan
Peninsula 2. – Singapore: Government Printer.
Ridley H. N. 1924: Flora of the Malay Peninsula 4. –
London: Reeve & Co.
Rudall P. J., Stevenson D. W. & Linder H. P. 1999:
Structure and systematics of Hanguana, a mono-
cotyledon of uncertain affinity. – Austr. Syst. Bot. 12:
311 – 330. [CrossRef]
Siti Nurfazilah A. R., Ahmad Sofiman O. & Boyce P.
C. [in press]: Studies on Hanguana (Commelinales-
Hanguanaceae) for Sunda I: Hanguana bakoensis, a
new forest species from Sarawak, Malaysian Borneo,
and notes on critical morphologies for elucidating
Hanguana taxonomy. – Acta Phytotax. Geobot.
Steenis-Kruseman M. J. van 2006: Cyclopedia of Male-
sian collectors. – Leiden: Nationaalherbarium Neder-
land; published at http://www.nationaalherbarium.nl/
FMcollectors/J/JackW.htm
Tillich H.-J. & Sill E. 1999: Systematische Studien zur
Morphologie und Anatomie von Hanguana Blume
(Hanguanaceae) und Flagellaria L. (Flagellariace­
ae), mit der Beschreibung einer neuen Art, Hanguana
bogneri spec. nov. – Sendtnera 6: 215 – 238.
Willdenowia 40 – 2010 213

Fig. 1. Hanguana exultans – A: flowering plant in habitat; B: infructescence, note the rather sharply ascending branches; C: detail
of the ventrally gibbose-ellipsoid fruits ripening pale yellow with a sessile stigma comprising 3 separate (not connate) orange
brown lobes; D: median leaf and inflorescence of H. pantiensis (left) compared with H. exultans (right). – A – D: Siti Nurfazilah &
al. HA-55; images © Rosazlina Bt Rusly.
214 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia

Fig. 2. Hanguana nitens – A: plants in habitat, note the arching (not stiffly erect) leaves; B: portion of a mature plant showing a
stolon and the conspicuous, long (2/3 – 1/4 of the leaf length) pseudopetiole; C: plicate leaf blade; D: portion of an immature infructes-
cence, compare the separate, erect, pointed stigma lobes with those of H. malayana (Fig. 7B); E: ripe fruits. – A – E: Siti Nurfazilah
& al. HA-48; images © Rosazlina Bt Rusly.
Willdenowia 40 – 2010 215

Fig. 3. Hanguana pantiensis – A: plant in habitat, note the dense panicle carried down in the leaves; B: detail of infructescence,
note the dense panicle with branches of the partial inflorescences ascending; C: ripe fruits, with the stigma obliquely to sublaterally
inserted by longitudinally bending of ovary; D: leaf bases showing litter-trapping. – A – D: Siti Nurfazilah & al. HA-56; images ©
Rosazlina Bt Rusly.
216 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia

Fig. 4. Hanguana podzolicola – A: plant in habitat showing the tall, leafless stem produced by older plants; B: detail of infructes-
cence, note the open nature of the panicle and the spreading branches of the partial inflorescences; C: ripe fruits, with the stigma
obliquely to sublaterally inserted by longitudinally bending of ovary; D: median leaf and inflorescence. – A – D: Siti Nurfazilah &
al. HA-49; images © Rosazlina Bt Rusly.
Willdenowia 40 – 2010 217

Fig. 5. Hanguana stenopoda – A – B: plants in habitat; B: detail of infructescence, note the partial inflorescences each with only 2
or rarely 3 branches and that the subtending bracts have fallen; D: ripe fruits with opaque tepals, briefly stipitate stigma, with the
lobes connate, deep chocolate brown, and the fruit with conspicuous black speckles. – A – D: Siti Nurfazilah & al. HA-60; images
© Siti Nurfazilah Bt Abdul Rahman.
218 Nurfazilah & al.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia

Fig. 6. Hanguana malayana – A: plants in habitat in Maludam N. P., Sarawak, note the stiffly erect leaves and the stolons floating
on the water surface; B: detail of semi-terrestrial stolons in habitat, Perak. – A: image © Mike Lo; B: image © Siti Nurfazilah Bt
Abdul Rahman.
Willdenowia 40 – 2010 219

Fig. 7. Hanguana malayana – A – B: female plants, note the short, robust branches, the green partial inflorescences, the ellipsoid
fruits, and the large sessile stigma almost obscuring the end of the fruit; C – D: male inflorescences, note the much longer, more
slender rather yellowish branches of the male inflorescence; after anthesis these branches droop prior to withering. – Images © Siti
Nurfazilah Bt Abdul Rahman.